Facultative nitrogen fixation by canopy legumes in a lowland tropical forest

Symbiotic dinitrogen (N₂) fixation is often invoked to explain the N richness of tropical forests as ostensibly N₂-fixing trees can be a major component of the community. Such arguments assume N₂ fixers are fixing N when present. However, in laboratory experiments, legumes consistently reduce N₂ fixation in response to increased soil N availability. These contrasting views of N₂ fixation as either obligate or facultative have drastically different implications for the N cycle of tropical forests. We tested these models by directly measuring N₂-fixing root nodules and nitrogenase activity of individual canopy-dominant legume trees (Inga sp.) across several lowland forest types. Fixation was substantial in disturbed forests and some gaps but near zero in the high N soils of mature forest. Our findings suggest that canopy legumes closely regulate N₂ fixation, leading to large variations in N inputs across the landscape, and low symbiotic fixation in mature forests despite abundant legumes.     

Effects of Increased Nitrogen Availability on C and N Cycles in Tropical Forests: A Meta-Analysis

Atmospheric N deposition is predicted to increase four times over its current status in tropical forests by 2030. Our ability to understand the effects of N enrichment on C and N cycles is being challenged by the large heterogeneity of the tropical forest biome. The specific response will depend on the forest’s nutrient status; however, few studies of N addition appear to incorporate the nutrient status in tropical forests, possibly due to difficulties in explaining how this status is maintained. We used a meta-analysis to explore the consequences of the N enrichment on C and N cycles in tropical montane and lowland forests. We tracked changes in aboveground and belowground plant C and N and in mineral soil in response to N addition. We found an increasing trend of plant biomass in montane forests, but not in lowland forests, as well as a greater increase in NO emission in montane forest compared with lowland forest. The N₂O and NO emission increase in both forest; however, the N₂O increase in lowland forest was significantly even at first time N addition. The NO emission increase showed be greater at first term compared with long term N addition. Moreover, the increase in total soil N, ammonium, microbial N, and dissolved N concentration under N enrichment indicates a rich N status of lowland forests. The available evidence of N addition experiments shows that the lowland forest is richer in N than montane forests. Finally, the greater increase in N leaching and N gas emission highlights the importance of study the N deposition effect on the global climate change.     

Throughfall exclusion and leaf litter addition drive higher rates of soil nitrous oxide emissions from a lowland wet tropical forest

Tropical forests are a significant global source of the greenhouse gas nitrous oxide (N₂O). Predicted environmental changes for this biome highlight the need to understand how simultaneous changes in precipitation and labile carbon (C) availability may affect soil N₂O production. We conducted a small‐scale throughfall and leaf litter manipulation in a lowland tropical forest in southwestern Costa Rica to test how potential changes in both water and litter derived labile C inputs to soils may alter N₂O emissions. Experimentally reducing throughfall in this wet tropical forest significantly increased soil emissions of N₂O, and our data suggest that at least part of this response was driven by an increase in the concentration of dissolved organic carbon [DOC] inputs delivered from litter to soil under the drier conditions. Furthermore, [DOC] was significantly correlated with N₂O emissions across both throughfall and litterfall manipulation plots, despite the fact that native NO₃^? pools in this site were generally small. Our results highlight the importance of understanding not only the potential direct effects of changing precipitation on soil biogeochemistry, but also the indirect effects resulting from interactions between the hydrologic, C and N cycles. Finally, over all sampling events we observed lower mean N₂O emissions (<1 ng N₂O‐N cm^?2 h^?1) than reported for many other lowland tropical forests, perhaps reflecting a more general pattern of increasing relative N constraints to biological activity as one moves from drier to wetter portions of the lowland tropical forest biome.     

Taxonomic identity determines N2 fixation by canopy trees across lowland tropical forests

Legumes capable of fixing atmospheric N₂ are abundant and diverse in many tropical forests, but the factors determining ecological patterns in fixation are unresolved. A long‐standing idea is that fixation depends on soil nutrients (N, P or Mo), but recent evidence shows that fixation may also differ among N₂‐fixing species. We sampled canopy‐height trees across five species and one species group of N₂‐fixers along a landscape P gradient, and manipulated P and Mo to seedlings in a shadehouse. Our results identify taxonomy as the major determinant of fixation, with P (and possibly Mo) only influencing fixation following tree‐fall disturbances. While 44% of trees did not fix N₂, other trees fixed at high rates, with two species functioning as superfixers across the landscape. Our results raise the possibility that fixation is determined by biodiversity, evolutionary history and species–specific traits (tree growth rate, canopy stature and response to disturbance) in the tropical biome.     

Nitrogen fixation does not axiomatically lead to phosphorus limitation in aquatic ecosystems

A long‐standing debate in ecology deals with the role of nitrogen and phosphorus in management and restoration of aquatic ecosystems. It has been argued that nutrient reduction strategies to combat blooms of phytoplankton or floating plants should solely focus on phosphorus (P). The underlying argument is that reducing nitrogen (N) inputs is ineffective because N₂‐fixing species will compensate for N deficits, thus perpetuating P limitation of primary production. A mechanistic understanding of this principle is, however, incomplete. Here, we use resource competition theory, a complex dynamic ecosystem model and a 32‐year field data set on eutrophic, floating‐plant dominated ecosystems to show that the growth of non‐N₂‐fixing species can become N limited under high P and low N inputs, even in the presence of N₂ fixing species. N₂‐fixers typically require higher P concentrations than non‐N₂‐fixers to persist. Hence, the N₂ fixers cannot deplete the P concentration enough for the non‐N₂‐fixing community to become P limited because they would be outcompeted. These findings provide a testable mechanistic basis for the need to consider the reduction of both N and P inputs to most effectively restore nutrient over‐enriched aquatic ecosystems.     

Nitrogen to phosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests

Increased atmospheric carbon dioxide (CO₂) concentrations and nitrogen (N) deposition induced by human activities have greatly influenced the stoichiometry of N and phosphorus (P). We used model forest ecosystems in open‐top chambers to study the effects of elevated CO₂ (ca. 700 μmol mol^?1) alone and together with N addition (100 kg N ha^?1 yr^?1) on N to P (N : P) ratios in leaves, stems and roots of five tree species, including four non‐N₂ fixers and one N₂ fixer, in subtropical China from 2006 to 2009. Elevated CO₂ decreased or had no effects on N : P ratios in plant tissues of tree species. N addition, especially under elevated CO₂, lowered N : P ratios in the N₂ fixer, and this effect was significant in the stems and the roots. However, only one species of the non‐N₂ fixers showed significantly lower N : P ratios under N addition in 2009, and the others were not affected by N addition. The reductions of N : P ratios in response to elevated CO₂ and N addition were mainly associated with the increases in P concentrations. Our results imply that elevated CO₂ and N addition could facilitate tree species to mitigate P limitation by more strongly influencing P dynamics than N in the subtropical forests.     

Immediate and long-term nitrogen oxide emissions from tropical forest soils exposed to elevated nitrogen input

Tropical nitrogen (N) deposition is projected to increase substantially within the coming decades. Increases in soil emissions of the climate‐relevant trace gases NO and N₂O are expected, but few studies address this possibility. We used N addition experiments to achieve N‐enriched conditions in contrasting montane and lowland forests and assessed changes in the timing and magnitude of soil N‐oxide emissions. We evaluated transitory effects, which occurred immediately after N addition, and long‐term effects measured at least 6 weeks after N addition. In the montane forest where stem growth was N limited, the first‐time N additions caused rapid increases in soil N‐oxide emissions. During the first 2 years of N addition, annual N‐oxide emissions were five times (transitory effect) and two times (long‐term effect) larger than controls. This contradicts the current assumption that N‐limited tropical montane forests will respond to N additions with only small and delayed increases in soil N‐oxide emissions. We attribute this fast and large response of soil N‐oxide emissions to the presence of an organic layer (a characteristic feature of this forest type) in which nitrification increased substantially following N addition. In the lowland forest where stem growth was neither N nor phosphorus (P) limited, the first‐time N additions caused only gradual and minimal increases in soil N‐oxide emissions. These first N additions were completed at the beginning of the wet season, and low soil water content may have limited nitrification. In contrast, the 9‐ and 10‐year N‐addition plots displayed instantaneous and large soil N‐oxide emissions. Annual N‐oxide emissions under chronic N addition were seven times (transitory effect) and four times (long‐term effect) larger than controls. Seasonal changes in soil water content also caused seasonal changes in soil N‐oxide emissions from the 9‐ and 10‐year N‐addition plots. This suggests that climate change scenarios, where rainfall quantity and seasonality change, will alter the relative importance of soil NO and N₂O emissions from tropical forests exposed to elevated N deposition.     

Substrate stoichiometry determines nitrogen fixation throughout succession in southern Chinese forests

The traditional view holds that biological nitrogen (N) fixation often peaks in early‐ or mid‐successional ecosystems and declines throughout succession based on the hypothesis that soil N richness and/or phosphorus (P) depletion become disadvantageous to N fixers. This view, however, fails to support the observation that N fixers can remain active in many old‐growth forests despite the presence of N‐rich and/or P‐limiting soils. Here, we found unexpected increases in N fixation rates in the soil, forest floor, and moss throughout three successional forests and along six age‐gradient forests in southern China. We further found that the variation in N fixation was controlled by substrate carbon(C) : N and C : (N : P) stoichiometry rather than by substrate N or P. Our findings highlight the utility of ecological stoichiometry in illuminating the mechanisms that couple forest succession and N cycling.     

Long‐term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil

Nitrogen (N) deposition is a component of global change that has considerable impact on belowground carbon (C) dynamics. Plant growth stimulation and alterations of fungal community composition and functions are the main mechanisms driving soil C gains following N deposition in N‐limited temperate forests. In N‐rich tropical forests, however, N deposition generally has minor effects on plant growth; consequently, C storage in soil may strongly depend on the microbial processes that drive litter and soil organic matter decomposition. Here, we investigated how microbial functions in old‐growth tropical forest soil responded to 13 years of N addition at four rates: 0 (Control), 50 (Low‐N), 100 (Medium‐N), and 150 (High‐N) kg N ha^?1 year^?1. Soil organic carbon (SOC) content increased under High‐N, corresponding to a 33% decrease in CO₂ efflux, and reductions in relative abundances of bacteria as well as genes responsible for cellulose and chitin degradation. A 113% increase in N₂O emission was positively correlated with soil acidification and an increase in the relative abundances of denitrification genes (narG and norB). Soil acidification induced by N addition decreased available P concentrations, and was associated with reductions in the relative abundance of phytase. The decreased relative abundance of bacteria and key functional gene groups for C degradation were related to slower SOC decomposition, indicating the key mechanisms driving SOC accumulation in the tropical forest soil subjected to High‐N addition. However, changes in microbial functional groups associated with N and P cycling led to coincidentally large increases in N₂O emissions, and exacerbated soil P deficiency. These two factors partially offset the perceived beneficial effects of N addition on SOC storage in tropical forest soils. These findings suggest a potential to incorporate microbial community and functions into Earth system models considering their effects on greenhouse gas emission, biogeochemical processes, and biodiversity of tropical ecosystems.     

Relationships among net primary productivity, nutrients and climate in tropical rain forest: a pan-tropical analysis

Tropical rain forests play a dominant role in global biosphere-atmosphere CO(2) exchange. Although climate and nutrient availability regulate net primary production (NPP) and decomposition in all terrestrial ecosystems, the nature and extent of such controls in tropical forests remain poorly resolved. We conducted a meta-analysis of carbon-nutrient-climate relationships in 113 sites across the tropical forest biome. Our analyses showed that mean annual temperature was the strongest predictor of aboveground NPP (ANPP) across all tropical forests, but this relationship was driven by distinct temperature differences between upland and lowland forests. Within lowland forests (相似文献   

Community structures of N2‐fixing bacteria associated with the phyllosphere of a Holm oak forest and their response to drought

Biological nitrogen (N) fixation is a key pathway in terrestrial ecosystems and is therefore critical for understanding the responses of ecosystems to global environmental changes. The free‐living diazotrophic community is distributed along the canopy‐to‐soil profile, but the ecological significance of epiphyllic N₂ fixers, despite their functional relevance, on plant foliar surfaces remains very poorly understood compared with the N₂‐fixing community in forest litter and soils. We assessed the community structure of N₂ fixers and overall bacteria by genetic fingerprinting (t‐RFLP) to explore the seasonal successional patterns of the microbial community in the natural phyllosphere of a Holm oak (Quercus ilex) forest submitted to 12‐year field experiment of rain exclusion mimicking the conditions of drought projected for the coming decades. Leaves of Holm oak were analysed in different seasons over a period of 1.5 years. The bacterial community of the phyllosphere did not correspond to the surrounding soil biome in the same area. These analyses provided field evidence for the presence of free‐living diazotrophs associated with the tissues of leaves of Holm oak, the dominant tree species of many Mediterranean forests. The results also revealed that the community composition is affected seasonally and inter‐annually by the environment, and that the composition shifts in response to climate change. Drought treatment increased the richness of the epiphyllic microbial community, especially during the summer. These changes were associated with higher C:N ratios of leaves observed in response to drought in semiarid areas. This epiphyllic microbiota that can potentially fix N₂ extends the capacity of plants to adapt to the environment.     

Distance‐dependent effects of soil‐derived biota on seedling survival of the tropical tree legume Ormosia semicastrata

Question: How do seed germination and subsequent seedling survival of O. semicastrata (Hance forma litchiifolia How) vary with respect to distance from parent trees and conspecific density in different types of tropical forest? Are there effects of soil biota on O. semicastrata that systematically depend on distance from parent trees and conspecific density? Do soil pathogens differently affect survival of O. semicastrata in different types of tropical forest? Location: Tropical lowland rain forest and tropical montane rain forest in Jianfengling National Nature Reserve, Hainan Island, China. Methods: Individual adult O. semicastrata trees were selected in lowland rain forest and montane rain forest. Soil was collected at a distance of 0‐5 m or 15‐20 m from the parent tree. Soil samples from each distance were combined into a bulk sample. Half of the soil sample was sterilized by autoclaving. Surface‐sterilized seeds were then added to the soil material in shade‐houses at both forests. Results: Germination of O. semicastrata seeds at low‐ or high‐seed density was barely affected by the sterilization procedure. In both forests, seedlings grown in non‐sterilized soil collected close to parent trees had significantly higher mortality compared to those in sterilized soil. In contrast, seedling survival with soil collected far from parent trees was not affected by the soil sterilization procedure. Conclusions: Host‐specific pathogens concentrated in the soil around parent trees may regulate community structure of tropical trees at the stage of seedling development.     

The response of heterotrophic activity and carbon cycling to nitrogen additions and warming in two tropical soils

Nitrogen (N) deposition is projected to increase significantly in tropical regions in the coming decades, where changes in climate are also expected. Additional N and warming each have the potential to alter soil carbon (C) storage via changes in microbial activity and decomposition, but little is known about the combined effects of these global change factors in tropical ecosystems. In this study, we used controlled laboratory incubations of soils from a long‐term N fertilization experiment to explore the sensitivity of soil C to increased N in two N‐rich tropical forests. We found that fertilization corresponded to significant increases in bulk soil C concentrations, and decreases in C loss via heterotrophic respiration (P< 0.05). The increase in soil C was not uniform among C pools, however. The active soil C pool decomposed faster with fertilization, while slowly cycling C pools had longer turnover times. These changes in soil C cycling with N additions corresponded to the responses of two groups of microbial extracellular enzymes. Smaller active C pools corresponded to increased hydrolytic enzyme activities; longer turnover times of the slowly cycling C pool corresponded to reduced activity of oxidative enzymes, which degrade more complex C compounds, in fertilized soils. Warming increased soil respiration overall, and N fertilization significantly increased the temperature sensitivity of slowly cycling C pools in both forests. In the lower elevation forest, respired CO₂ from fertilized cores had significantly higher Δ¹⁴C values than control soils, indicating losses of relatively older soil C. These results indicate that soil C storage is sensitive to both N deposition and warming in N‐rich tropical soils, with interacting effects of these two global change factors. N deposition has the potential to increase total soil C stocks in tropical forests, but the long‐term stability of this added C will likely depend on future changes in temperature.     

Nitrogen deposition contributes to soil acidification in tropical ecosystems

Elevated anthropogenic nitrogen (N) deposition has greatly altered terrestrial ecosystem functioning, threatening ecosystem health via acidification and eutrophication in temperate and boreal forests across the northern hemisphere. However, response of forest soil acidification to N deposition has been less studied in humid tropics compared to other forest types. This study was designed to explore impacts of long‐term N deposition on soil acidification processes in tropical forests. We have established a long‐term N‐deposition experiment in an N‐rich lowland tropical forest of Southern China since 2002 with N addition as NH₄NO₃ of 0, 50, 100 and 150 kg N ha^?1 yr^?1. We measured soil acidification status and element leaching in soil drainage solution after 6‐year N addition. Results showed that our study site has been experiencing serious soil acidification and was quite acid‐sensitive showing high acidification (pH_(H2O)<4.0), negative water‐extracted acid neutralizing capacity (ANC) and low base saturation (BS,< 8%) throughout soil profiles. Long‐term N addition significantly accelerated soil acidification, leading to depleted base cations and decreased BS, and further lowered ANC. However, N addition did not alter exchangeable Al³⁺, but increased cation exchange capacity (CEC). Nitrogen addition‐induced increase in SOC is suggested to contribute to both higher CEC and lower pH. We further found that increased N addition greatly decreased soil solution pH at 20 cm depth, but not at 40 cm. Furthermore, there was no evidence that Al³⁺ was leaching out from the deeper soils. These unique responses in tropical climate likely resulted from: exchangeable H⁺ dominating changes of soil cation pool, an exhausted base cation pool, N‐addition stimulating SOC production, and N saturation. Our results suggest that long‐term N addition can contribute measurably to soil acidification, and that shortage of Ca and Mg should receive more attention than soil exchangeable Al in tropical forests with elevated N deposition in the future.     

Relationships among precipitation regime, nutrient availability, and carbon turnover in tropical rain forests

The effect of high precipitation regime in tropical forests is poorly known despite indications of its potentially negative effects on nutrient availability and carbon (C) cycling. Our goal was to determine if there was an effect of high rainfall on nitrogen (N) and phosphorous (P) availability and indexes of C cycling in lowland tropical rain forests exposed to a broad range of mean annual precipitation (MAP). We predicted that C turnover time would increase with MAP while the availability of N and P would decrease. We studied seven Neotropical lowland forests covering a MAP range between 2,700 and 9,500 mm. We used radiocarbon (?¹⁴C) from the atmosphere and respired from soil organic matter to estimate residence time of C in plants and soils. We also used C, N, and P concentrations and the stable isotope ratio of N (δ¹⁵N) in live and dead plant tissues and in soils as proxies for nutrient availability. Negative δ¹⁵N values indicated that the wettest forests had N cycles that did not exhibit isotope-fractionating losses and were potentially N-limited. Element ratios (N:P and C:P) in senescent leaves, litter, and live roots showed that P resorption increased considerably with MAP, which points towards increasing P-limitation under high MAP regimes. Soil C content increased with MAP but C turnover time only showed a weak relationship with MAP, probably due to variations in soil parent material and age along the MAP gradient. In contrast, comparing C turnover directly to nutrient availability showed strong relationships between C turnover time, N availability (δ¹⁵N), and P availability (N:P) in senescent leaves and litter. Thus, an effect of MAP on carbon cycling appeared to be indirectly mediated by nutrient availability. Our results suggest that soil nutrient availability plays a central role in the dynamic of C cycling in tropical rain forests.     

Soil fertility and fine root dynamics in response to 4 years of nutrient (N,P, K) fertilization in a lowland tropical moist forest,Panama

The question of how tropical trees cope with infertile soils has been challenging to address, in part, because fine root dynamics must be studied in situ. We used annual fertilization with nitrogen (N as urea, 12.5 g N m^?2 year^?1), phosphorus (P as superphosphate, 5 g P m^?2 year^?1) and potassium (K as KCl, 5 g K m^?2 year^?1) within 38 ha of old‐growth lowland tropical moist forest in Panama and examined fine root dynamics with minirhizotron images. We expected that added P, above all, would (i) decrease fine root biomass but, (ii) have no impact on fine root turnover. Soil in the study area was moderately acidic (pH = 5.28), had moderate concentrations of exchangeable base cations (13.4 cmol kg^?1), low concentrations of Bray‐extractable phosphate (PO₄ = 2.2 mg kg^?1), and modest concentrations of KCl‐extractable nitrate (NO₃ = 5.0 mg kg^?1) and KCl‐extractable ammonium (NH₄ = 15.5 mg kg^?1). Added N increased concentrations of KCl‐extractable NO₃ and acidified the soil by one pH unit. Added P increased concentrations of Bray‐extractable PO₄ and P in the labile fraction. Concentrations of exchangeable K were elevated in K addition plots but reduced by N additions. Fine root dynamics responded to added K rather than added P. After 2 years, added K decreased fine root biomass from 330 to 275 g m^?2. The turnover coefficient of fine roots <1 mm diameter ranged from 2.6 to 4.4 per year, and the largest values occurred in plots with added K. This study supported the view that biomass and dynamics of fine roots respond to soil nutrient availability in species‐rich, lowland tropical moist forest. However, K rather than P elicited root responses. Fine roots smaller than 1 mm have a short lifetime (<140 days), and control of fine root production by nutrient availability in tropical forests deserves more study.     

Mechanism of application nursery cultivation arbuscular mycorrhizal seedling in watermelon in the field

Arbuscular mycorrhizal fungi (AMF) colonisation of plant root facilitates the absorption of nutrients such as phosphorus (P) and enhances plant biotic and abiotic resistance generally. However, arbuscular mycorrhiza (AM) colonisation decreases with application of chemical fertiliser. Here, we investigated whether AMF inoculation in nurseries would facilitate AM colonisation and take physiological and ecological functions in watermelon (Citrullus lanatus) in the field. Pot experiments were carried out to study the change of AMF colonised seedling on physiology and gene expression in nursery site. Field experiments were performed to investigate the effect of nursery AMF inoculation on yield, quality and disease resistance of watermelon in the field. The results showed that nursery‐inoculated seedlings produced more dry matter and root surface area than non‐inoculated seedlings. Expression of the secretory purple acid phosphatase (PAP) genes ClaPAP10 and ClaPAP26 was up‐regulated following AMF colonisation. Accordingly, acid phosphatase activities at the root surface and P concentrations in seedling were enhanced. After transplantation to the field, the shoot dry matter and P concentration in old stem were higher in the nursery AMF inoculated seedlings than that in non‐AMF inoculated seedling. AMF inoculation also induced increase of yields and decrease of wilt disease indexes and soluble sugar content. In addition, acid phosphatase activities and AMF spore densities were increased by nursery‐inoculation in watermelon rhizosphere soil in the field. In conclusion, nursery colonisation AMF seedling enhanced watermelon growth and yield by improving the root growth and P acquisition in nursery cultivating stage, as well as optimised soil properties in the field. Nursery cultivation of watermelon seedling with AMF was an effective technique to reduce wilt disease in continuous cropped management in watermelon.     

The response of microbial biomass and hydrolytic enzymes to a decade of nitrogen, phosphorus, and potassium addition in a lowland tropical rain forest

Nutrient availability is widely considered to constrain primary productivity in lowland tropical forests, yet there is little comparable information for the soil microbial biomass. We assessed microbial nutrient limitation by quantifying soil microbial biomass and hydrolytic enzyme activities in a long-term nutrient addition experiment in lowland tropical rain forest in central Panama. Multiple measurements were made over an annual cycle in plots that had received a decade of nitrogen, phosphorus, potassium, and micronutrient addition. Phosphorus addition increased soil microbial carbon (13 %), nitrogen (21 %), and phosphorus (49 %), decreased phosphatase activity by ~65 % and N-acetyl β-glucosaminidase activity by 24 %, but did not affect β-glucosidase activity. In contrast, addition of nitrogen, potassium, or micronutrients did not significantly affect microbial biomass or the activity of any enzyme. Microbial nutrients and hydrolytic enzyme activities all declined markedly in the dry season, with the change in microbial biomass equivalent to or greater than the annual nutrient flux in fine litter fall. Although multiple nutrients limit tree productivity at this site, we conclude that phosphorus limits microbial biomass in this strongly-weathered lowland tropical forest soil. This finding indicates that efforts to include enzymes in biogeochemical models must account for the disproportionate microbial investment in phosphorus acquisition in strongly-weathered soils.     

Partitioning of soil phosphorus among arbuscular and ectomycorrhizal trees in tropical and subtropical forests

Partitioning of soil phosphorus (P) pools has been proposed as a key mechanism maintaining plant diversity, but experimental support is lacking. Here, we provided different chemical forms of P to 15 tree species with contrasting root symbiotic relationships to investigate plant P acquisition in both tropical and subtropical forests. Both ectomycorrhizal (ECM) and arbuscular mycorrhizal (AM) trees responded positively to addition of inorganic P, but strikingly, ECM trees acquired more P from a complex organic form (phytic acid). Most ECM tree species and all AM tree species also showed some capacity to take up simple organic P (monophosphate). Mycorrhizal colonisation was negatively correlated with soil extractable P concentration, suggesting that mycorrhizal fungi may regulate organic P acquisition among tree species. Our results support the hypothesis that ECM and AM plants partition soil P sources, which may play an ecologically important role in promoting species coexistence in tropical and subtropical forests.     

Variability of indices of macronutrient availability in soils at different spatial scales along an elevation transect in tropical moist forests (NE Ecuador)

The availability of key plant nutrients may change with elevation in tropical mountains due to altitudinal gradients in temperature and moisture which affect pedogenesis and nutrient cycling. In a transect from upper lowland to montane forests in NE Ecuador, we tested the hypotheses that (1) the availability of P is low in low-elevation forests but increases upslope, while the availability of N is relatively high at low elevations but decreases with elevation, and (2) increasing amounts of calcium, magnesium and potassium are stored on top of the soil with progressive humus accumulation toward higher elevations, likely to improve nutrient availability. In each 20 plots in undisturbed natural forest at 500, 1000, 1500 and 2000 m?a.s.l., we measured in situ N net mineralization and nitrification rate (N_NM and N_NI, buried bag method), plant-available phosphorus (P_a, resin-bag method), and salt-exchangeable calcium, potassium and magnesium concentrations (Ca_ex, K_ex, Mg_ex) in the organic and mineral topsoils. N_NM and N_NI, and the Ca_ex, K_ex and Mg_ex concentrations were much more variable at the plot level than across the four elevations, while P_a varied equally at small and large spatial scales. P_a increased more than 10fold from 500 to 2000 m. The net release of nitrate dominated over ammonium at all elevations. While mass-related N_NM and N_NI rates and also organic matter C/N ratio in the topsoil remained invariant along the slope, N_NM and N_NI rates per ground area decreased by about 40% from 500 to 2000 m. Thus, the N_NM/P_a ratio decreased markedly with elevation proving our first hypothesis. In support of the second hypothesis, the pools of Ca_ex, Mg_ex and K_ex in the organic layers increased with elevation, demonstrating the key role that organic topsoil horizons are playing for forest nutrition at high elevations. We suggest that the large difference in N versus P availability of tropical (upper) lowland and montane forests is likely to be a key factor influencing the species composition and productivity along tropical mountain slopes.